Degenerate neutrality creates evolvable fitness landscapes

Understanding how systems can be designed to be evolvable is fundamental to research in optimization, evolution, and complex systems science. Many researchers have thus recognized the importance of evolvability, i.e. the ability to find new variants of higher fitness, in the fields of biological evolution and evolutionary computation. Recent studies by Ciliberti et al (Proc. Nat. Acad. Sci., 2007) and Wagner (Proc. R. Soc. B., 2008) propose a potentially important link between the robustness and the evolvability of a system. In particular, it has been suggested that robustness may actually lead to the emergence of evolvability. Here we study two design principles, redundancy and degeneracy, for achieving robustness and we show that they have a dramatically different impact on the evolvability of the system. In particular, purely redundant systems are found to have very little evolvability while systems with degeneracy, i.e. distributed robustness, can be orders of magnitude more evolvable. These results offer insights into the general principles for achieving evolvability and may prove to be an important step forward in the pursuit of evolvable representations in evolutionary computation

Degeneracy: a link between evolvability, robustness and complexity in biological systems

A full accounting of biological robustness remains elusive; both in terms of the mechanisms by which robustness is achieved and the forces that have caused robustness to grow over evolutionary time. Although its importance to topics such as ecosystem services and resilience is well recognized, the broader relationship between robustness and evolution is only starting to be fully appreciated. A renewed interest in this relationship has been prompted by evidence that mutational robustness can play a positive role in the discovery of adaptive innovations (evolvability) and evidence of an intimate relationship between robustness and complexity in biology. This paper offers a new perspective on the mechanics of evolution and the origins of complexity, robustness, and evolvability. Here we explore the hypothesis that degeneracy, a partial overlap in the functioning of multi-functional components, plays a central role in the evolution and robustness of complex forms. In support of this hypothesis, we present evidence that degeneracy is a fundamental source of robustness, it is intimately tied to multi-scaled complexity, and it establishes conditions that are necessary for system evolvability

Degeneracy: a design principle for achieving robustness and evolvability

Robustness, the insensitivity of some of a biological system's functionalities to a set of distinct conditions, is intimately linked to fitness. Recent studies suggest that it may also play a vital role in enabling the evolution of species. Increasing robustness, so is proposed, can lead to the emergence of evolvability if evolution proceeds over a neutral network that extends far throughout the fitness landscape. Here, we show that the design principles used to achieve robustness dramatically influence whether robustness leads to evolvability. In simulation experiments, we find that purely redundant systems have remarkably low evolvability while degenerate, i.e. partially redundant, systems tend to be orders of magnitude more evolvable. Surprisingly, the magnitude of observed variation in evolvability can neither be explained by differences in the size nor the topology of the neutral networks. This suggests that degeneracy, a ubiquitous characteristic in biological systems, may be an important enabler of natural evolution. More generally, our study provides valuable new clues about the origin of innovations in complex adaptive systems.Comment: Accepted in the Journal of Theoretical Biology (Nov 2009

Visualising the Global Structure of Search Landscapes: Genetic Improvement as a Case Study

The search landscape is a common metaphor to describe the structure of computational search spaces. Different landscape metrics can be computed and used to predict search difficulty. Yet, the metaphor falls short in visualisation terms because it is hard to represent complex landscapes, both in terms of size and dimensionality. This paper combines Local Optima Networks, as a compact representation of the global structure of a search space, and dimensionality reduction, using the t-Distributed Stochastic Neighbour Embedding (t-SNE) algorithm, in order to both bring the metaphor to life and convey new insight into the search process. As a case study, two benchmark programs, under a Genetic Improvement bug-fixing scenario, are analysed and visualised using the proposed method. Local Optima Networks for both iterated local search and a hybrid genetic algorithm, across different neighbourhoods, are compared, highlighting the differences in how the landscape is explored

Networked buffering: a basic mechanism for distributed robustness in complex adaptive systems

A generic mechanism - networked buffering - is proposed for the generation of robust traits in complex systems. It requires two basic conditions to be satisfied: 1) agents are versatile enough to perform more than one single functional role within a system and 2) agents are degenerate, i.e. there exists partial overlap in the functional capabilities of agents. Given these prerequisites, degenerate systems can readily produce a distributed systemic response to local perturbations. Reciprocally, excess resources related to a single function can indirectly support multiple unrelated functions within a degenerate system. In models of genome:proteome mappings for which localized decision-making and modularity of genetic functions are assumed, we verify that such distributed compensatory effects cause enhanced robustness of system traits. The conditions needed for networked buffering to occur are neither demanding nor rare, supporting the conjecture that degeneracy may fundamentally underpin distributed robustness within several biotic and abiotic systems. For instance, networked buffering offers new insights into systems engineering and planning activities that occur under high uncertainty. It may also help explain recent developments in understanding the origins of resilience within complex ecosystems. \ud \u

Dynamics of transcription factor binding site evolution

Evolution of gene regulation is crucial for our understanding of the phenotypic differences between species, populations and individuals. Sequence-specific binding of transcription factors to the regulatory regions on the DNA is a key regulatory mechanism that determines gene expression and hence heritable phenotypic variation. We use a biophysical model for directional selection on gene expression to estimate the rates of gain and loss of transcription factor binding sites (TFBS) in finite populations under both point and insertion/deletion mutations. Our results show that these rates are typically slow for a single TFBS in an isolated DNA region, unless the selection is extremely strong. These rates decrease drastically with increasing TFBS length or increasingly specific protein-DNA interactions, making the evolution of sites longer than ~10 bp unlikely on typical eukaryotic speciation timescales. Similarly, evolution converges to the stationary distribution of binding sequences very slowly, making the equilibrium assumption questionable. The availability of longer regulatory sequences in which multiple binding sites can evolve simultaneously, the presence of "pre-sites" or partially decayed old sites in the initial sequence, and biophysical cooperativity between transcription factors, can all facilitate gain of TFBS and reconcile theoretical calculations with timescales inferred from comparative genetics.Comment: 28 pages, 15 figure

Neutral Networks of Real-World Programs and their Application to Automated Software Evolution

The existing software development ecosystem is the product of evolutionary forces, and consequently real-world software is amenable to improvement through automated evolutionary techniques. This dissertation presents empirical evidence that software is inherently robust to small randomized program transformations, or \u27mutations. Simple and general mutation operations are demonstrated that can be applied to software source code, compiled assembler code, or directly to binary executables. These mutations often generate variants of working programs that differ significantly from the original, yet remain fully functional. Applying successive mutations to the same software program uncovers large \u27neutral networks\u27 of fully functional variants of real-world software projects. These properties of \u27mutational robustness\u27 and the corresponding \u27neutral networks\u27 have been studied extensively in biology and are believed to be related to the capacity for unsupervised evolution and adaptation. As in biological systems, mutational robustness and neutral networks in software systems enable automated evolution. The dissertation presents several applications that leverage software neutral networks to automate common software development and maintenance tasks. Neutral networks are explored to generate diverse implementations of software for improving runtime security and for proactively repairing latent bugs. Next, a technique is introduced for automatically repairing bugs in the assembler and executables compiled from off-the-shelf software. As demonstration, a proprietary executable is manipulated to patch security vulnerabilities without access to source code or any aid from the software vendor. Finally, software neutral networks are leveraged to optimize complex nonfunctional runtime properties. This optimization technique is used to reduce the energy consumption of the popular PARSEC benchmark applications by 20% as compared to the best available public domain compiler optimizations. The applications presented herein apply evolutionary computation techniques to existing software using common software engineering tools. By enabling evolutionary techniques within the existing software development toolchain, this work is more likely to be of practical benefit to the developers and maintainers of real-world software systems

Evolvability-guided Optimization of Linear Deformation Setups for Evolutionary Design Optimization

Richter A. Evolvability-guided Optimization of Linear Deformation Setups for Evolutionary Design Optimization. Bielefeld: Universität Bielefeld; 2019.Andreas Richter gratefully acknowledges the financial support from Honda Research Institute Europe (HRI-EU).This thesis targets efficient solutions for optimal representation setups for evolutionary design optimization problems. The representation maps the abstract parameters of an optimizer to a meaningful variation of the design model, e.g., the shape of a car. Thereby, it determines the convergence speed to and the quality of the final result. Thus, engineers are eager to employ well-tuned representations to achieve high-quality design solutions. But, setting up optimal representations is a cumbersome process because the setup procedure requires detailed knowledge about the objective functions, e.g., a fluid dynamics simulation, and the parameters of the employed representation itself. Thus, we target efficient routines to set up representations automatically to support engineers from their tedious, partly manual work. Inspired by the concept of evolvability, we present novel quality criteria for the evaluation of linear deformations commonly applied as representations. We define and analyze the criteria variability, regularity, and improvement potential which measure the expected quality and convergence speed of an evolutionary design optimization process based on the linear deformation setup. Moreover, we target the efficient optimization of deformation setups with respect to these three criteria. In dynamic design optimization scenarios a suitable compromise between exploration and exploitation is crucial for efficient solutions. We discuss the construction of optimal compromises for these dynamic scenarios with our criteria because they characterize exploration and exploitation. As a result an engineer can initialize and adjust the deformation setup for improved convergence speed of a design process and for enhanced quality of the design solutions with our methods

How codon choice determines evolvability and evolutionary robustness in short linear motifs

Short linear motifs, made up of 2-10 amino acids in linear sequence space, are a central component of cellular decision making through proteins. They form a modular system in cells where combinations of domains and motifs are used as basic functional building blocks through interactions. Functions mediated through these motifs include cellular localisation, post-translational modifications, degradation and general protein-protein interactions. Since motifs are made up of a small number of amino acids they have unusual evolutionary properties, for instance they can evolve de novo, or be lost, through a small number of substitutions. This is of particular importance in pathogens such as viruses. Many viruses evolve new host-like motifs to interact with the host and change the regulation and signalling landscape within host cells to mediate infection. In this body of work, I have used influenza as a model to elucidate aspects of the evolutionary properties of motifs. I have been able to leverage recent progress made in determining nucleotide mutation rates and have developed a model for motif evolution that is defined from the nucleotide and codon levels. Simulations using this methodology suggested that different codons have varying propensities to evolve into amino acids within a linear motif. In other words, some sequences have higher motif evolvability. The simulations also indicated a fitness benefit to use some codons over others to encode linear motifs, due to the varying propensity to evolve. These findings suggest that motifs that are encoded by specific codons have higher motif evolutionary robustness, i.e. they can tolerate more mutations without affecting function. I went on to investigate if these predicted properties have played a role in motif evolution in influenza. I found that conserved motifs in influenza use the codons inferred to have higher evolutionary robustness. This would lead to increased fitness, as motifs are less often lost through mutations. I also found that this mutational robustness acts on stop codon usage in influenza, suggesting an explanation for an old observation of predominant use of TAA in many organisms. Interestingly, it also appears that evolutionary robustness of a motif can be varied to tune the rate of motif change, which influenza utilises in glycosylation motifs that interface with the host immune system. Finally, I investigated whether the codon choice and evolvability at early stages of viral host shifts could be used to predict the emergence of functional motifs. I have found that motif evolvability can aid the prediction of motif emergence. For influenza strains H1N1 and H3N2, which were introduced in the human population from birds during the 1900s, the sequence of the early strains could be used to predict the majority of the glycosylation sites that would emerge the following decades. The predictability of motif emergence could have important implications for vaccination efforts. The methodologies developed here, and the observations made about how motif evolution is shaped by codon choices in a predictable way will be important for a better understanding of the evolution of complexity and regulation involving motifs. This may have implications for complex diseases such as cancers, and for our understanding of the evolution of pathogen innovations and functionality